Novel gene involved in brassinosteroid responses

ABSTRACT

There is provided a polynucleotide encoding a plant gene capable of controlling a signal transduction system for brassinosteroid hormone, the polynucleotide encoding an amino acid sequence from Met at position 1 to Arg at position 1057 of SEQ ID NO: 2 in the SEQUENCE LISTING, including any polynucleotide encoding an amino acid sequence in which one or more amino acids are deleted, substituted or added to the amino acid sequence.

BACKGROUND OF THE INVENTION

[0001] 1. Field of the Invention

[0002] The present invention relates to a novel gene. In particular, thepresent invention relates to a novel gene in plants which encodes aprotein having the function of controlling an in-vivo signaltransduction system in a physiological reaction system againstbrassinosteroid hormone.

[0003] 2. Description of the Related Art

[0004] Transposons are mutagenic genes which are known to be ubiquitousin animal, yeast, bacterial, and plant genomes. Transposons areclassified into two classes, Class I and Class II, depending on theirtransposition mechanisms. Transposons belonging to Class II aretransposed in the form of DNAs without being replicated. Known Class IItransposons include the Ac/Ds, Spm/dSpm and Mu elements of Zea mays(Fedoroff, 1989, Cell 56, 181-191: Fedoroff et al., 1983, Cell 35,235-242; Schiefelbein et al., 1985, Proc. Natl. Acad. Sci. USA 82,4783-4787), and the Tam element of Antirrhinum majus (Bonas et al.,1984, EMBO J., 3, 1015-1019). Class II transposons are widely used forgene isolation techniques which utilize transposon tagging. Suchtechniques utilize the fact that a transposon induces physiological andmorphological changes when inserted into genes. The affected gene can beisolated by detecting such changes (Bancroft et al., 1993, The PlantCell, 5, 631-638; Colasanti et al., 1998, Cell, 93, 593-603: Gray etal., 1997, Cell, 89, 25-31; Keddie et al., 1998, The Plant Cell, 10,877-887: Whitham et al., 1994, Cell, 78, 1101-1115).

[0005] Transposons belonging to Class I, also referred to asretrotransposons, are replicated and transposed via RNA intermediates.Class I transposons were first identified and characterized inDrosophila and in yeasts. However, recent studies have revealed thatClass I transposons are ubiquitous in plant genomes and account for asubstantial portion of the genomes (Bennetzen, 1996, Trends Microbiolo.,4, 347-353; Voytas, 1996, Science, 274, 737-738). A large majority ofretrotransposons appear to be inactive. Recent studies indicate thatsome of these retrotransposons are activated under stress conditionssuch as injuries, pathogenic attacks, or cell culture (Grandbastien,1998, Trends in Plant Science, 3, 181-187; Wessler, 1996, Curr. Biol. 6,959-961; Wessler et al., 1995, Curr. Opin. Genet. Devel. 5, 814-821).Activation under stress conditions has been reported for Tnt1 A and Tto1in tobacco (Pouteau et al., 1994, Plant J., 5, 535-542: Takeda et al.,1988, Plant Mol. Biol., 36, 365-376), and Tos17 in rice (Hirochika etal., 1996, Proc. Natl. Acad. Sci. USA, 93, 7783-7788), for example.

[0006] The Tos17 retrotransposon of rice is one of the most-extensivelystudied plant Class I elements in plants. Tosl7 was cloned by an RT-PCRmethod using a degenerate primer prepared based on a conservative aminoacid sequence in reverse transcription enzyme domains between Ty1-copiaretroelements (Hirochika et al., 1992, Mol. Gen. Genet., 233, 209-216).Tos17 is 4.3 kb long, and has two 138 bp LTRs (long chain terminalrepetitions) and PBS (primer binding sites) complementary to the 3′ endof the start methionine tRNA (Hirochika et al., 1996, supra). Tos17transcription is strongly activated through tissue culture, and its copynumber increases with culture time. In Nipponbare, a model Japonicacultivar used for genome analysis, two copies of Tos17 are initiallypresent, which are increased to 5 to 30 copies in a regenerated plantafter tissue culture (Hirochika et al., 1996, supra). Unlike Class IItransposons which were characterized in yeasts and Drosophila, Tos17 istransposed in chromosomes in random manners and causes stable mutation,and therefore provides a powerful tool for functional analysis of ricegenes (Hirochika, 1997, Plant Mol. Biol. 35, 231-240; 1999, MolecularBiology of. Rice (ed. by K. Shimamoto, Springer-Verlag, 43-58).

SUMMARY OF THE INVENTION

[0007] The present invention relates to a polynucleotide encoding aplant gene capable of controlling a signal transduction system forbrassinosteroid hormone, the polynucleotide encoding an amino acidsequence from Met at position 1 to Arg at position 1057 of SEQ ID NO: 2in the SEQUENCE LISTING, including any polynucleotide encoding an aminoacid sequence in which one or more amino acids are deleted, substitutedor added to the amino acid sequence.

[0008] In one embodiment of the invention, the polynucleotide may bederived from rice.

[0009] In another embodiment of the invention, the polynucleotide may beas represented by SEQ ID NO: 1 in the SEQUENCE LISTING.

[0010] The present invention further relates to methods for controllingvarious effects in plants in which brassinosteroid hormone is involved,e.g., growth promotion, yield increase, quality improvement, maturationenhancement, and tolerance against biotic and abiotic stresses.

[0011] The inventors diligently conducted systematic analyses ofphenotypes of plants having a newly transposed To1 7 copy and sequencesadjoining Tos17 target sites with respect to rice. As a result, theinventors found a dwarf rice mutation obtained from Tos17 insertion, andisolated the gene responsible for this mutation by utilizing Tos17 as atag, thereby accomplishing the present invention.

[0012] Thus, the invention described herein makes possible the advantageof providing a novel plant gene which can be provided by using Tos17.

[0013] This and other advantages of the present invention will becomeapparent to those skilled in the art upon reading and understanding thefollowing detailed description with reference to the accompanyingfigures.

BRIEF DESCRIPTION OF THE DRAWINGS

[0014]FIGS. 1A and 1B are photographs showing a brassinosteroidnon-sensitive mutant having Tos17 inserted therein, which was foundamong regenerated Akitakomachi lineage. On the left of each figure isshown a brassinosteroid non-sensitive mutant having Tos17 insertedtherein. On the right of each figure is shown a wild type plant body.FIG. 1A evidences an influence toward dwarfism and upright form. FIG. 1Bevidences an influence toward malformation of grain hulls.

[0015]FIGS. 2A and 2B are photographs showing a brassinosteroidnon-sensitive mutant having Tos17 inserted therein, which was foundamong regenerated Nipponbare lineage. On the left of each figure isshown a brassinosteroid non-sensitive mutant having Tos17 insertedtherein. On the right of each figure is shown a wild type plant body.FIG. 2A evidences an influence toward dwarfism and upright form. FIG. 2Bevidences an influence toward malformation of grain hulls.

[0016]FIG. 3A shows a Northern analysis autoradiogram of RNA extractedfrom the leaves of a brassinosteroid non-sensitive mutant (Akitakomachi)and RNA extracted from various organs of a wild type rice plant(Nipponbare).

[0017]FIG. 3B shows a Northern analysis autoradiogram of RNA extractedfrom brassinosteroid non-sensitive mutants and RNA extracted from wildtype rice plants. The left-hand side of FIG. 3B shows a comparisonbetween wild types and mutants obtained by using a 5′ probe. Theright-hand side of FIG. 3B shows a comparison between wild types andmutants obtained by using a 3′ probe.

[0018]FIG. 4 shows an amino acid sequence of the novel rice gene whichcontrols a physiological reaction system induced by brassinosteroidhormone, together with characteristic sequences found therein (wherenuclear localization signals and an ATP/GTP binding motif can beobserved).

[0019]FIG. 5A shows a brassinosteroid leaf blade bending experimentationusing a mutated line (A0369) derived from Akitakomachi. The left-handside shows results of wild type plants, whereas the right-hand sideshows results of mutants.

[0020]FIG. 5B shows a brassinosteroid leaf blade bending experimentationusing a mutated line (NC6148) derived from Nipponbare. The left-handside shows results of wild type plants, whereas the right-hand sideshows results of mutants.

DESCRIPTION OF THE PREFERRED EMBODIMENTS

[0021] The present invention provides a method for producing an improvedplant, the method utilizing a novel plant gene which can be provided byusing Tos17.

[0022] According to the present invention, there is provided apolynucleotide encoding a plant gene capable of controlling variouseffects in which brassinosteroid hormone is involved. As used herein,the term “capable of controlling various effects” means the ability tocontrol various effects in plants in which brassinosteroid hormone isinvolved, e.g., growth promotion, yield increase, quality improvement,maturation enhancement, and tolerance against biotic and abioticstresses, including dwarfism, upright form, and malformation of grainhulls, thereby providing a number of agriculturally useful effects asare attained by treatments with brassinosteroid hormone agriculturalchemicals. The term “plants” encompasses both monocotyledons anddicotyledons.

[0023] A polynucleotide encoding a plant gene capable of controlling asignal transduction system for brassinosteroid hormone according to thepresent invention is, for example, a polynucleotide encoding an aminoacid sequence from Met at position 1 to Arg at position 1057 of SEQ IDNO: 2 in the SEQUENCE LISTING, including any polynucleotide encoding anamino acid sequence in which one or more amino acids are deleted,substituted or added to the aforementioned amino acid sequence.

[0024] A polynucleotide encoding a plant gene capable of controllingvarious effects in which brassinosteroid hormone is involved encompassesany polynucleotides which have at least about 80% sequence homology,preferably at least about 85% sequence homology, and more preferably atleast about 90% sequence homology, still more preferably at least about95% sequence homology, and most preferably at least about 99% sequencehomology, with an amino acid sequence from Met at position 1 to Arg atposition 1057 of SEQ ID NO: 2 in the SEQUENCE LISTING, so long as theyare capable of controlling various effects in plants in whichbrassinosteroid hormone is involved. The term “sequence homology”indicates a degree of identity between two polynucleotide sequences tobe compared with each other. The rate (%) of sequence homology betweentwo polynucleotide sequences for comparison is calculated by, afteroptimally aligning the two polynucleotide sequences for comparison,obtaining a matched position number indicating the number of positionsat which identical (“matched”) nucleic acid bases (e.g., A, T, C, G, U,or I) are present in both sequences, dividing the matched positionnumber by total number of bases in the polynucleotide sequences forcomparison, and multiplying the quotient by 100. The sequence homologycan be calculated by using the following sequencing tools, for example:a Unix base program designated GCG Wisconsin Package (Program Manual forthe Wisconsin Package, Version 8, September 1994, Genetics ComputerGroup, 575 Science Drive Madison, Wis., USA 53711; Rice, P. (1996)Program Manual for EGCG Package, Peter Rice, The Sanger Centre, HinxtonHall, Cambridge, CB10 1RQ, England), and the ExPASy World Wide Webmolecular biology server (Geneva University Hospital and University ofGeneva, Geneva, Switzerland).

[0025] Cells into which genes have been introduced are first selectedbased on drug resistance, e.g., hygromycin resistance, and thenregenerated into plant bodies by using usual methods.

[0026] The terminology and laboratory procedures described throughoutthe present specification are directed to those which are well-known andcommonly employed in the art. Standard techniques may be used forrecombination methods, polynucleotide synthesis, microorganismsculturing, and transformation (e.g., electroporation). Such techniquesand procedures are generally known from various standard textbooksavailable in the field or by way of the present specification (includinga generally-referenced textbook by Sambrook et al., Molecular Cloning: ALaboratory Manual, 2nd edition (1989) Cold Spring Harbor LaboratoryPress, Cold Spring Harbor, N.Y.). Such literature is incorporated hereinby reference.

[0027] The polynucleotide according to the present invention can beobtained by using the method described herein, for example. However, thepolynucleotide according to the present invention may also be obtainedby any chemical synthesis process based on the sequence disclosedherein. For example, the polynucleotide according to the presentinvention may be synthesized by using a polynucleotide synthesizeravailable from Applied Bio Systems in accordance with the instructionsprovided by the manufacturer.

[0028] Methods of PCR amplification are well-known in the art (PCRTechnology: Principles and Applications for DNA Amplification, ed. H AErlich, Freeman Press, NewYork, N.Y. (1992); PCR Protocols: A Guide toMethods and Applications, Innis, Gelfland, Snisky, and White, AcademicPress, San Diego, Calif. (1990); Mattila et al. (1991) Nucleic AcidsRes. 19: 4967; Eckert, K. A. and Kunkel, T. A. (1991) PCR Methods andApplications 1: 17; PCR, McPherson, Quirkes, and Taylor, IRL Press,Oxford). Such literature is incorporated herein by reference.

EXAMPLES

[0029] Hereinafter, the present invention will be described by way ofexamples which are of illustrative but non-limiting nature.

Example 1

[0030] Activation of Tos17 Via Culture

[0031] Using fully ripened seeds of Nipponbare and Akitakomachi, whichare varieties of Japonica subspecies, induction of calli and cellsuspension culture were carried out as described earlier (Hirochika etal., 1996, supra). The activation of Tos17 which was used for genedestruction was carried out following the method of Ohtsuki (1990) (riceprotoplast culture system, Food and Agricultural Research DevelopmentAssociation). In summary, fully ripened seeds of rice were cultured inan MS medium having 2,4-dichlorophenoxyacetic acid (2,4-D) added thereto(2 mg/ml) (Ohtsuki (1990), supra) (25° C., 1 month), to induce callusformation. The resultant calluses were cultured for 5 months in an N6liquid medium having 2,4-D added thereto (Ohtsuki (1990), supra), andthereafter placed on a redifferentiation medium (Ohtsuki (1990), supra),whereby redifferentiated rice plants were obtained (first generation(R1) plants).

Example 2

[0032] Isolation of sequences adjoining Tos17

[0033] Utilizing each of the regenerated R1 rice plants obtainedaccording to Example 1 as a first strain, about 1000 R1 seeds werecollected from each strain and grown on a paddy field to obtain secondgeneration (R2) plants, which were subjected to a morphologicalanalysis. As a result of observing the phenotypes of the respectiveplant bodies in the R2 group, it was learned that about ¼ of the R2group of an Akitakomachi strain A0369 exhibit the “dwarfism, uprightform, and malformation of grain hulls” phenotype (FIGS. 1A and 1B). Inthe regenerated group of Akitakomachi, dwarfism, upright form, andmalformation of grain hulls were observed for brassinosteroidinsensitive mutants (FIG. 1A, left, and FIG. 1B, left), as compared withthe wild type (FIG. 1A, right, and FIG. 1B, right). The isolation ofadjacent sequences of transposed Tos17, which is co-segragating with thephenotypes, was carried out by an IPCR method (Ochman et al., GeneticsNov; 120(3): 621-3(1988) and Triglia et al., Nucleic Acids Res Aug,25:(16): 8186(1988)) The total DNA of A0369 was digested with XbaI, and aligation process was performed in a large quantity of solution, therebyobtaining self-ligated circular molecules. In the self-ligated circularmolecules, the adjacent sequences are flanking the internal sequence ofTos17. As a result, amplification was successfully carried out by usualPCR methods using an outward primer pair (T17TAIL3:GAGAGCATCATCGGTTACATCTTCTC; T17-1950R: TCTAGCAGTCTCAATGATGTGGCG) basedon the known sequence of Tos17.

Example 3

[0034] Search for Alleles

[0035] Using the sequence obtained according to Example 2, lineage inwhich Tos17 had been inserted at a different site in the same gene wasselected by PCR from the regenerated rice group of Nipponbare. As aresult, a line (NC6148) which similarly exhibited dwarfism, uprightform, and malformation of grain hulls were observed for brassinosteroidwas selected. That is, in the regenerated rice group of Nipponbare, aswell, dwarfism, upright form, and malformation of grain hulls wereobserved for brassinosteroid insensitive mutants (FIG. 2A, left, andFIG. 2B, left), relative to the wild type (FIG. 2A, right, and FIG. 2B,right). It was concluded that these common mutations were results of thesame gene having been destroyed.

Example 4

[0036] Expression and Analysis of the Causative Gene

[0037] From the group of R2 rice plants (selfed progeny from the A0369and NC6148 strains) obtained according to Examples 2 and 3, individualsexhibiting mutation were identified from normal individuals. RNA wasprepared from both groups of individuals, and the expression specificitywas analyzed through Northern analysis. After agarose electrophoresis,the RNA obtained from individuals exhibiting mutation and the RNAobtained from normal individuals were allowed to adsorb to nylonmembranes. DNA fragments which were obtained by amplifying via PCR asequence (positions 5775-6638 of the genomic sequence) on the 5′ sideand a sequence (positions 8175-8765 of the genomic sequence) on the 3′side of the Tos17 insertion site in both mutated lines were labeled with³²P-dCTP. By using these as probes, expression specificity was analyzedthrough Northern analysis (FIGS. 3A and 3B). As seen from the Northernanalysis autoradiogram shown in FIG. 3A, a band (about 4.3 kb) indicatedby an arrow was confirmed to be expressed in all observed organs of thewild type. However, in the mutants, transcripts of abnormal sizes wereobserved due to the insertion of Tos17, indicating that the naturalfunction of the wild type is lost (FIG. 3B).

Example 5

[0038] Structural Analysis of the Causative Gene

[0039] Using the sequence obtained according to Example 2 as a probe,the corresponding cDNA and genomic clone were obtained from a cDNAlibrary and a genomic library. Their structures are shown in SEQ ID Nos:1 and 3. It was learned that this gene includes 6 exons and 5 introns,encoding 1057 amino acids, and that Tos17 had been inserted at the 4thand 5th exons in two mutants, respectively. Moreover, motif searchresults suggested the presence of nuclear localization signal 1 (aminoacid residues 329-367 of SEQ ID NO: 2, Robbins & Dingwall consensussequence; a search result by PSORT program(http://psort.ims.u-tokyo.ac.jp/)) and nuclear localization signal 2(amino acid residues 457-460, 595-600 of SEQ ID NO: 2, 4 amino acidnuclear localization pattern signal; a search result by PSORT program(http://psort.ims.u-tokyo.ac.jp/)) as well as the presence of an ATP/GTPbinding domain (amino acid residues 526-533 of SEQ ID NO: 2; a searchresult by a motif search service onGenomenet(http://www.genome.ad.jp/)). Thus, the possibility of this genebeing involved in signal transduction was suggested (FIG. 4).

Example 6

[0040] Brassinosteroid Sensitivity Evaluation

[0041] The present gene was deduced to be a factor involved in thesignal transduction system for plant hormones, taking note of the factsthat the present gene was expressed in all plant bodies and thatpleiotropic influences resulted from destroying this gene, as well asthe possibility that the gene might be a factor involved in the signaltransduction system. Presuming that the signal transduction system isthat for brassinosteroid hormone in view of the resultant upright form,the inventors performed a leaf blade bend response test as abrassinosteroid response test, by using brassinolide, which is one kindof brassinosteroid hormone. The second leaf of rice which was allowed togerminate in the dark was cut off, and immersed in a 1 ng/ml ofbrassinosteroid solution for 48 hours. The wild type individuals havingthe wild type genes showed bending of the leaf blades and leaf sheathjunctions (left-hand side in FIGS. 5A and 5B), showing response tobrassinolide, whereas mutant individuals showed little bending thereof(right-hand side in FIGS. 5A and 5B), indicating that the destruction ofthe present gene resulted in the loss of response to brassinosteroid.From the above results, it was revealed that the present gene is a geneinvolved in the signal transduction system for brassinosteroid hormone.

[0042] The above examples are illustrative, and by no means limiting, ofvarious aspects of the present invention and the manners in which theoligonucleotide according to the present invention can be made andutilized.

[0043] Thus, according to the present invention, a novel polynucleotideis provided which is capable of controlling various effects in whichbrassinosteroid hormone is involved, the polynucleotide being of use inplant breeding. By introducing the present polynucleotide into plantsand artificially controlling various effects in which brassinosteroidhormone is involved, it is expected that effects such as growthpromotion, yield increase, quality improvement, maturation enhancement,and tolerance against biotic and abiotic stresses can be controlled,thereby providing a number of agriculturally useful effects as areattained by treatments with brassinosteroid hormone agriculturalchemicals.

[0044] Various other modifications will be apparent to and can bereadily made by those skilled in the art without departing from thescope and spirit of this invention. Accordingly, it is not intended thatthe scope of the claims appended hereto be limited to the description asset forth herein, but rather that the claims be broadly construed.

1 3 1 4310 DNA Oryza sativa CDS (655)..(3828) 1 ctcttctcca ctccaaatcccttcttaccc tattcccctc cccccgcagc ttctcttcct 60 cctgcagtac tcgccgccaccaccaccgcg ccgccgccgc cggccgcgtt ccgagaccca 120 ctcgatcgga atccaccgcggcgcgcccgc gcgcctgcgt cctcttcctt ccccgggagc 180 cgaccgacca cggcgaccagtcgatctccc tctccgggcg ccaaccgcgt cttagcttca 240 tcgaatccac cgccccaccccgcatctcct cctcctcctc cgacgacgac gactactact 300 agtcttctcc aataagcccccctcccgctc cccccgcctg aagaagaagc agcagctagc 360 tccggggaga ggtcgacggcgcgccgggta gatcgcgccc cgccccgcct gcgtcgcggc 420 tgtcggagca aacgcaaaccccccaggttg ttctagcgtg tgcagcggct agctgattga 480 ttgtcttctg tgatatatccagagctcgtg ttttgtggtt tgtggtttgt ggtttgtgct 540 tggattgttg atgtgctaattcgcggcgtt acaagatcac tgctggattg atattgagtt 600 gtgcctcggc tgtgctagctgtgtgttgat tctctcctcg tcgtggtgat cgat atg 657 Met 1 gag att gtt gca gtagat cag gag gga gct cgt gtt gtt ggg acg aac 705 Glu Ile Val Ala Val AspGln Glu Gly Ala Arg Val Val Gly Thr Asn 5 10 15 tgt atg ctt gct cgt ggtgga act ggt gct gta gcg cca gtg ttg gag 753 Cys Met Leu Ala Arg Gly GlyThr Gly Ala Val Ala Pro Val Leu Glu 20 25 30 ctg aca gcg acg cct cgt caggat gca gcc gct gaa gct ggt gta gac 801 Leu Thr Ala Thr Pro Arg Gln AspAla Ala Ala Glu Ala Gly Val Asp 35 40 45 gaa ccg gca caa cac caa tgc gagcat ttc tcc ata aga ggg tat gtt 849 Glu Pro Ala Gln His Gln Cys Glu HisPhe Ser Ile Arg Gly Tyr Val 50 55 60 65 gct ctt ctt cag aag aag gat ccaaaa ttc tgc tct cta tct cgg att 897 Ala Leu Leu Gln Lys Lys Asp Pro LysPhe Cys Ser Leu Ser Arg Ile 70 75 80 ttc cat gac cag aaa aaa tgt gat gaacac aaa gct agt tca agc cca 945 Phe His Asp Gln Lys Lys Cys Asp Glu HisLys Ala Ser Ser Ser Pro 85 90 95 ttt tct gta gca aag ttt cga cga tgg gattgc tcg aag tgc ttg gat 993 Phe Ser Val Ala Lys Phe Arg Arg Trp Asp CysSer Lys Cys Leu Asp 100 105 110 aag ttg aaa act tca gat aat gga aca gcacca aga act ctt ccc gca 1041 Lys Leu Lys Thr Ser Asp Asn Gly Thr Ala ProArg Thr Leu Pro Ala 115 120 125 aag cag aat ggc aca agt gat ggt tgc tccatc aca ttt gtt cgg agc 1089 Lys Gln Asn Gly Thr Ser Asp Gly Cys Ser IleThr Phe Val Arg Ser 130 135 140 145 act ttt gtg cct gct agt gtt ggt tcccaa aaa gtg tct cct agc aca 1137 Thr Phe Val Pro Ala Ser Val Gly Ser GlnLys Val Ser Pro Ser Thr 150 155 160 caa tca tct caa ggg aag aat gct gataga tca act ctt cca aag agt 1185 Gln Ser Ser Gln Gly Lys Asn Ala Asp ArgSer Thr Leu Pro Lys Ser 165 170 175 gtg caa gaa ggc aat gac tcc aaa tgcaat gcg cct tct ggc aag aat 1233 Val Gln Glu Gly Asn Asp Ser Lys Cys AsnAla Pro Ser Gly Lys Asn 180 185 190 gga gct gct gag gcc aat act gat tcacca atg aaa gat ttg caa ggg 1281 Gly Ala Ala Glu Ala Asn Thr Asp Ser ProMet Lys Asp Leu Gln Gly 195 200 205 cca gcc caa aat tat gat gtg gca gcaaat gtc tct gag gac aac act 1329 Pro Ala Gln Asn Tyr Asp Val Ala Ala AsnVal Ser Glu Asp Asn Thr 210 215 220 225 tct gtt gat gtt ggg gct tta cctgaa gtt ccc cag att aca tgg cac 1377 Ser Val Asp Val Gly Ala Leu Pro GluVal Pro Gln Ile Thr Trp His 230 235 240 ata gaa gta aat ggt gca gat caacct cca tcc act cca aaa ctt tct 1425 Ile Glu Val Asn Gly Ala Asp Gln ProPro Ser Thr Pro Lys Leu Ser 245 250 255 gaa gtg gtc ctc aaa aga aat gaagat gaa aat gga aaa act gaa gag 1473 Glu Val Val Leu Lys Arg Asn Glu AspGlu Asn Gly Lys Thr Glu Glu 260 265 270 act ctt gtt gct gag cag tgc aatttg acc aaa gat cct aac cca atg 1521 Thr Leu Val Ala Glu Gln Cys Asn LeuThr Lys Asp Pro Asn Pro Met 275 280 285 tct gga aag gaa cgt gat cag gttgct gag cag tgc aat ttg acc aaa 1569 Ser Gly Lys Glu Arg Asp Gln Val AlaGlu Gln Cys Asn Leu Thr Lys 290 295 300 305 gat ccg aaa cca gtg tct gggcag aaa tgt gag cag atc tgc aat gag 1617 Asp Pro Lys Pro Val Ser Gly GlnLys Cys Glu Gln Ile Cys Asn Glu 310 315 320 cca tgt gaa gaa gtt gtt ctcaaa aga agc tcc aaa tct aag agg aag 1665 Pro Cys Glu Glu Val Val Leu LysArg Ser Ser Lys Ser Lys Arg Lys 325 330 335 acg gat aag aag ttg atg aagaag cag cag cac agc aag aaa cgc act 1713 Thr Asp Lys Lys Leu Met Lys LysGln Gln His Ser Lys Lys Arg Thr 340 345 350 gcc cag gct gat gtt tca gatgca aag ctt tgt cgg aga aag cca aaa 1761 Ala Gln Ala Asp Val Ser Asp AlaLys Leu Cys Arg Arg Lys Pro Lys 355 360 365 aag gtg cgg ctt cta tca gaaatt ata aat gct aac cag gtt gag gat 1809 Lys Val Arg Leu Leu Ser Glu IleIle Asn Ala Asn Gln Val Glu Asp 370 375 380 385 tct aga agt gac gaa gttcat cgt gaa aat gcc gct gat ccc tgt gag 1857 Ser Arg Ser Asp Glu Val HisArg Glu Asn Ala Ala Asp Pro Cys Glu 390 395 400 gat gat aga agt acc atcccg gtc ccg atg gaa gta agc atg gat att 1905 Asp Asp Arg Ser Thr Ile ProVal Pro Met Glu Val Ser Met Asp Ile 405 410 415 cct gtt agc aac cat acagtg gga gaa gat ggg tta aaa tca agt aag 1953 Pro Val Ser Asn His Thr ValGly Glu Asp Gly Leu Lys Ser Ser Lys 420 425 430 aac aag aca aaa cgc aaatac tct gat gtt gta gat gat gga tca tca 2001 Asn Lys Thr Lys Arg Lys TyrSer Asp Val Val Asp Asp Gly Ser Ser 435 440 445 ctt atg aac tgg ctg aatgga aaa aag aaa aga act gga agt gtg cat 2049 Leu Met Asn Trp Leu Asn GlyLys Lys Lys Arg Thr Gly Ser Val His 450 455 460 465 cac aca gtt gct catcca gct ggg aat ttg agc aac aaa aaa gtg aca 2097 His Thr Val Ala His ProAla Gly Asn Leu Ser Asn Lys Lys Val Thr 470 475 480 ccc act gcg agt actcag cat gat gat gag aat gat act gaa aat ggt 2145 Pro Thr Ala Ser Thr GlnHis Asp Asp Glu Asn Asp Thr Glu Asn Gly 485 490 495 ctt gac aca aat atgcat aag aca gat gtc tgt cag cat gta tca gaa 2193 Leu Asp Thr Asn Met HisLys Thr Asp Val Cys Gln His Val Ser Glu 500 505 510 atc tcc aca cag aggtgc tca tca aag ggg aaa aca gcg ggt ttg agt 2241 Ile Ser Thr Gln Arg CysSer Ser Lys Gly Lys Thr Ala Gly Leu Ser 515 520 525 aag ggg aaa aca cattca gct gct agt acc aaa tat ggt ggt gaa agc 2289 Lys Gly Lys Thr His SerAla Ala Ser Thr Lys Tyr Gly Gly Glu Ser 530 535 540 545 acc aga aat ggtcag aac ata cat gta ctc agc gca gaa gat caa tgc 2337 Thr Arg Asn Gly GlnAsn Ile His Val Leu Ser Ala Glu Asp Gln Cys 550 555 560 cag atg gaa accgaa aac tct gtt ctg agt cac tcg gca aag gtt tct 2385 Gln Met Glu Thr GluAsn Ser Val Leu Ser His Ser Ala Lys Val Ser 565 570 575 cca gct gag catgat atc caa att atg tct gac ctt cat gag cag agt 2433 Pro Ala Glu His AspIle Gln Ile Met Ser Asp Leu His Glu Gln Ser 580 585 590 cta ccc aag aagaaa aag aag caa aaa ctt gaa gtg act cgt gaa aaa 2481 Leu Pro Lys Lys LysLys Lys Gln Lys Leu Glu Val Thr Arg Glu Lys 595 600 605 cag acc atg atagat gac atc ccc atg gat att gtt gaa ctg cta gct 2529 Gln Thr Met Ile AspAsp Ile Pro Met Asp Ile Val Glu Leu Leu Ala 610 615 620 625 aaa aac cagcat gag agg cag ctt atg act gag act gat tgt tct gac 2577 Lys Asn Gln HisGlu Arg Gln Leu Met Thr Glu Thr Asp Cys Ser Asp 630 635 640 atc aac cgtatt caa tcc aag aca act gct gat gat gat tgt gta ata 2625 Ile Asn Arg IleGln Ser Lys Thr Thr Ala Asp Asp Asp Cys Val Ile 645 650 655 gta gct gccaag gat ggt tca gat tat gca tca agt gtg ttt gac act 2673 Val Ala Ala LysAsp Gly Ser Asp Tyr Ala Ser Ser Val Phe Asp Thr 660 665 670 aat tcc caacag aag tcc ttg gca tcc caa agt aca cag aag gag tta 2721 Asn Ser Gln GlnLys Ser Leu Ala Ser Gln Ser Thr Gln Lys Glu Leu 675 680 685 cag ggt catttg gca ttg acc aca caa gag tct cca cat cct cag aac 2769 Gln Gly His LeuAla Leu Thr Thr Gln Glu Ser Pro His Pro Gln Asn 690 695 700 705 ttt cagtct act cag gaa cag cag aca cat ttg cgg atg gaa gaa atg 2817 Phe Gln SerThr Gln Glu Gln Gln Thr His Leu Arg Met Glu Glu Met 710 715 720 gtc actatt gct gca agc tca cca cta ttt tca cat cat gat gat cag 2865 Val Thr IleAla Ala Ser Ser Pro Leu Phe Ser His His Asp Asp Gln 725 730 735 tat attgct gaa gca cca act gaa cat tgg ggc cgt aag gac gca aag 2913 Tyr Ile AlaGlu Ala Pro Thr Glu His Trp Gly Arg Lys Asp Ala Lys 740 745 750 aag ctaacg tgg gag caa ttt aag gcc act aca aga aat tct cca gca 2961 Lys Leu ThrTrp Glu Gln Phe Lys Ala Thr Thr Arg Asn Ser Pro Ala 755 760 765 gca acatgt ggt gct caa ttt aga cct ggt atc caa gca gtt gac ttg 3009 Ala Thr CysGly Ala Gln Phe Arg Pro Gly Ile Gln Ala Val Asp Leu 770 775 780 785 acttct act cat gtc atg gga tct tcc agc aat tat gca tct cgc caa 3057 Thr SerThr His Val Met Gly Ser Ser Ser Asn Tyr Ala Ser Arg Gln 790 795 800 ccagta att gcg cca ctg gac cgc tat gct gaa aga gcg gtt aac cag 3105 Pro ValIle Ala Pro Leu Asp Arg Tyr Ala Glu Arg Ala Val Asn Gln 805 810 815 gtccat gca aga aat ttt cca agc aca ata gca acc atg gaa gcg agt 3153 Val HisAla Arg Asn Phe Pro Ser Thr Ile Ala Thr Met Glu Ala Ser 820 825 830 aagtta tgt gat cgg aga aat gct gga caa gta gtc ttg tat cct aaa 3201 Lys LeuCys Asp Arg Arg Asn Ala Gly Gln Val Val Leu Tyr Pro Lys 835 840 845 gaatcc atg cct gcg acg cat ctt ctg aga atg atg gat cca tca aca 3249 Glu SerMet Pro Ala Thr His Leu Leu Arg Met Met Asp Pro Ser Thr 850 855 860 865tta gca agc ttc ccc aac tat gga act tct agc agg aac cag atg gag 3297 LeuAla Ser Phe Pro Asn Tyr Gly Thr Ser Ser Arg Asn Gln Met Glu 870 875 880tct caa ctt cat aat tct cag tat gca cat aat cag tac aaa gga tca 3345 SerGln Leu His Asn Ser Gln Tyr Ala His Asn Gln Tyr Lys Gly Ser 885 890 895acc agc aca tca tat ggc agt aac ctg aat gga aag att cca ttg aca 3393 ThrSer Thr Ser Tyr Gly Ser Asn Leu Asn Gly Lys Ile Pro Leu Thr 900 905 910ttc gaa gac tta tca cgg cat cag ctg cat gat ctg cac aga cct tta 3441 PheGlu Asp Leu Ser Arg His Gln Leu His Asp Leu His Arg Pro Leu 915 920 925cgc cca cat cct aga gtt ggt gtg ctt ggc tcc ttg ctg cag aag gaa 3489 ArgPro His Pro Arg Val Gly Val Leu Gly Ser Leu Leu Gln Lys Glu 930 935 940945 att gca aac tgg tcg gag aac tgt ggc aca caa tct ggt tat aag tta 3537Ile Ala Asn Trp Ser Glu Asn Cys Gly Thr Gln Ser Gly Tyr Lys Leu 950 955960 gga gtg tca aca gga ata aca tcg cat cag atg aac aga aag gaa cat 3585Gly Val Ser Thr Gly Ile Thr Ser His Gln Met Asn Arg Lys Glu His 965 970975 ttt gaa gcc ctg aat tct gga atg ttt tca gca aaa tgg aat gca ttg 3633Phe Glu Ala Leu Asn Ser Gly Met Phe Ser Ala Lys Trp Asn Ala Leu 980 985990 cag ttg ggt tct gtt agc tcc agt gca gat ttt tta tca gcg agg aac 3681Gln Leu Gly Ser Val Ser Ser Ser Ala Asp Phe Leu Ser Ala Arg Asn 995 10001005 agc ata gct caa tct tgg acc aga ggc aag ggt aaa atg gtt cat ccc3729 Ser Ile Ala Gln Ser Trp Thr Arg Gly Lys Gly Lys Met Val His Pro1010 1015 1020 1025 ttg gat cgg ttt gtg aga cag gat atc tgt ata act aacaag aac cca 3777 Leu Asp Arg Phe Val Arg Gln Asp Ile Cys Ile Thr Asn LysAsn Pro 1030 1035 1040 gct gat ttt act aca atc agt aac gat aac gag tatatg gat tac cgc 3825 Ala Asp Phe Thr Thr Ile Ser Asn Asp Asn Glu Tyr MetAsp Tyr Arg 1045 1050 1055 tga agcagaaagt ggtgtgcata attcctgaacatttacaatc atacatttca 3878 tctttatggc gccaaatagt catactgtaa gaggagggctttgctggatc tgctgtaagg 3938 cttcttgtaa gttgtggatg ccccattttc tggatgggaacctgccagac agtgaacaag 3998 ggctttgcaa ggtgcagcat ccggtttttg ttttgccagtccaagaaacg tcctcctgtt 4058 actttgtagt tgtactcata ctagtgcgct tgtttgtacaaggagaaatg tgtaaccttg 4118 ttgaaaaaat gtctccccca ttttgtaatt accataaggaggtttatagt gttgtgagct 4178 gtgtgtgact gacggcgaga aatggttttg tcggtgttaaggttgaaacg actagctctc 4238 gttatcaatg tgttgtaaac ttctagattg atgtgttaccttactcttga agtcaacacc 4298 ggagaattta ca 4310 2 1057 PRT Oryza sativa 2Met Glu Ile Val Ala Val Asp Gln Glu Gly Ala Arg Val Val Gly Thr 1 5 1015 Asn Cys Met Leu Ala Arg Gly Gly Thr Gly Ala Val Ala Pro Val Leu 20 2530 Glu Leu Thr Ala Thr Pro Arg Gln Asp Ala Ala Ala Glu Ala Gly Val 35 4045 Asp Glu Pro Ala Gln His Gln Cys Glu His Phe Ser Ile Arg Gly Tyr 50 5560 Val Ala Leu Leu Gln Lys Lys Asp Pro Lys Phe Cys Ser Leu Ser Arg 65 7075 80 Ile Phe His Asp Gln Lys Lys Cys Asp Glu His Lys Ala Ser Ser Ser 8590 95 Pro Phe Ser Val Ala Lys Phe Arg Arg Trp Asp Cys Ser Lys Cys Leu100 105 110 Asp Lys Leu Lys Thr Ser Asp Asn Gly Thr Ala Pro Arg Thr LeuPro 115 120 125 Ala Lys Gln Asn Gly Thr Ser Asp Gly Cys Ser Ile Thr PheVal Arg 130 135 140 Ser Thr Phe Val Pro Ala Ser Val Gly Ser Gln Lys ValSer Pro Ser 145 150 155 160 Thr Gln Ser Ser Gln Gly Lys Asn Ala Asp ArgSer Thr Leu Pro Lys 165 170 175 Ser Val Gln Glu Gly Asn Asp Ser Lys CysAsn Ala Pro Ser Gly Lys 180 185 190 Asn Gly Ala Ala Glu Ala Asn Thr AspSer Pro Met Lys Asp Leu Gln 195 200 205 Gly Pro Ala Gln Asn Tyr Asp ValAla Ala Asn Val Ser Glu Asp Asn 210 215 220 Thr Ser Val Asp Val Gly AlaLeu Pro Glu Val Pro Gln Ile Thr Trp 225 230 235 240 His Ile Glu Val AsnGly Ala Asp Gln Pro Pro Ser Thr Pro Lys Leu 245 250 255 Ser Glu Val ValLeu Lys Arg Asn Glu Asp Glu Asn Gly Lys Thr Glu 260 265 270 Glu Thr LeuVal Ala Glu Gln Cys Asn Leu Thr Lys Asp Pro Asn Pro 275 280 285 Met SerGly Lys Glu Arg Asp Gln Val Ala Glu Gln Cys Asn Leu Thr 290 295 300 LysAsp Pro Lys Pro Val Ser Gly Gln Lys Cys Glu Gln Ile Cys Asn 305 310 315320 Glu Pro Cys Glu Glu Val Val Leu Lys Arg Ser Ser Lys Ser Lys Arg 325330 335 Lys Thr Asp Lys Lys Leu Met Lys Lys Gln Gln His Ser Lys Lys Arg340 345 350 Thr Ala Gln Ala Asp Val Ser Asp Ala Lys Leu Cys Arg Arg LysPro 355 360 365 Lys Lys Val Arg Leu Leu Ser Glu Ile Ile Asn Ala Asn GlnVal Glu 370 375 380 Asp Ser Arg Ser Asp Glu Val His Arg Glu Asn Ala AlaAsp Pro Cys 385 390 395 400 Glu Asp Asp Arg Ser Thr Ile Pro Val Pro MetGlu Val Ser Met Asp 405 410 415 Ile Pro Val Ser Asn His Thr Val Gly GluAsp Gly Leu Lys Ser Ser 420 425 430 Lys Asn Lys Thr Lys Arg Lys Tyr SerAsp Val Val Asp Asp Gly Ser 435 440 445 Ser Leu Met Asn Trp Leu Asn GlyLys Lys Lys Arg Thr Gly Ser Val 450 455 460 His His Thr Val Ala His ProAla Gly Asn Leu Ser Asn Lys Lys Val 465 470 475 480 Thr Pro Thr Ala SerThr Gln His Asp Asp Glu Asn Asp Thr Glu Asn 485 490 495 Gly Leu Asp ThrAsn Met His Lys Thr Asp Val Cys Gln His Val Ser 500 505 510 Glu Ile SerThr Gln Arg Cys Ser Ser Lys Gly Lys Thr Ala Gly Leu 515 520 525 Ser LysGly Lys Thr His Ser Ala Ala Ser Thr Lys Tyr Gly Gly Glu 530 535 540 SerThr Arg Asn Gly Gln Asn Ile His Val Leu Ser Ala Glu Asp Gln 545 550 555560 Cys Gln Met Glu Thr Glu Asn Ser Val Leu Ser His Ser Ala Lys Val 565570 575 Ser Pro Ala Glu His Asp Ile Gln Ile Met Ser Asp Leu His Glu Gln580 585 590 Ser Leu Pro Lys Lys Lys Lys Lys Gln Lys Leu Glu Val Thr ArgGlu 595 600 605 Lys Gln Thr Met Ile Asp Asp Ile Pro Met Asp Ile Val GluLeu Leu 610 615 620 Ala Lys Asn Gln His Glu Arg Gln Leu Met Thr Glu ThrAsp Cys Ser 625 630 635 640 Asp Ile Asn Arg Ile Gln Ser Lys Thr Thr AlaAsp Asp Asp Cys Val 645 650 655 Ile Val Ala Ala Lys Asp Gly Ser Asp TyrAla Ser Ser Val Phe Asp 660 665 670 Thr Asn Ser Gln Gln Lys Ser Leu AlaSer Gln Ser Thr Gln Lys Glu 675 680 685 Leu Gln Gly His Leu Ala Leu ThrThr Gln Glu Ser Pro His Pro Gln 690 695 700 Asn Phe Gln Ser Thr Gln GluGln Gln Thr His Leu Arg Met Glu Glu 705 710 715 720 Met Val Thr Ile AlaAla Ser Ser Pro Leu Phe Ser His His Asp Asp 725 730 735 Gln Tyr Ile AlaGlu Ala Pro Thr Glu His Trp Gly Arg Lys Asp Ala 740 745 750 Lys Lys LeuThr Trp Glu Gln Phe Lys Ala Thr Thr Arg Asn Ser Pro 755 760 765 Ala AlaThr Cys Gly Ala Gln Phe Arg Pro Gly Ile Gln Ala Val Asp 770 775 780 LeuThr Ser Thr His Val Met Gly Ser Ser Ser Asn Tyr Ala Ser Arg 785 790 795800 Gln Pro Val Ile Ala Pro Leu Asp Arg Tyr Ala Glu Arg Ala Val Asn 805810 815 Gln Val His Ala Arg Asn Phe Pro Ser Thr Ile Ala Thr Met Glu Ala820 825 830 Ser Lys Leu Cys Asp Arg Arg Asn Ala Gly Gln Val Val Leu TyrPro 835 840 845 Lys Glu Ser Met Pro Ala Thr His Leu Leu Arg Met Met AspPro Ser 850 855 860 Thr Leu Ala Ser Phe Pro Asn Tyr Gly Thr Ser Ser ArgAsn Gln Met 865 870 875 880 Glu Ser Gln Leu His Asn Ser Gln Tyr Ala HisAsn Gln Tyr Lys Gly 885 890 895 Ser Thr Ser Thr Ser Tyr Gly Ser Asn LeuAsn Gly Lys Ile Pro Leu 900 905 910 Thr Phe Glu Asp Leu Ser Arg His GlnLeu His Asp Leu His Arg Pro 915 920 925 Leu Arg Pro His Pro Arg Val GlyVal Leu Gly Ser Leu Leu Gln Lys 930 935 940 Glu Ile Ala Asn Trp Ser GluAsn Cys Gly Thr Gln Ser Gly Tyr Lys 945 950 955 960 Leu Gly Val Ser ThrGly Ile Thr Ser His Gln Met Asn Arg Lys Glu 965 970 975 His Phe Glu AlaLeu Asn Ser Gly Met Phe Ser Ala Lys Trp Asn Ala 980 985 990 Leu Gln LeuGly Ser Val Ser Ser Ser Ala Asp Phe Leu Ser Ala Arg 995 1000 1005 AsnSer Ile Ala Gln Ser Trp Thr Arg Gly Lys Gly Lys Met Val His 1010 10151020 Pro Leu Asp Arg Phe Val Arg Gln Asp Ile Cys Ile Thr Asn Lys Asn1025 1030 1035 1040 Pro Ala Asp Phe Thr Thr Ile Ser Asn Asp Asn Glu TyrMet Asp Tyr 1045 1050 1055 Arg 3 9455 DNA Oryza sativa 3 gatctatcgttgattggatt tcgctgggct agctaggtag acaagggttc aaaatgtgac 60 gaacattattgagctaataa ataacgagaa accaacttgg tatataggtc aatttcaaaa 120 gaaacaagctgacaaaattc gtccaatttc actagttttt gtcagtaatt gaatggcaat 180 catggttatcgacaaaaccg cttaggagtg ctatttggtg atggaatggt ttataaaact 240 ttggaccggagtagcagtac aatggcttgt ctgaacaggc taggtagcat agtaggtcct 300 tttgccttggttgcactgtt ctgtcggctt ataggggaat ctattggctt aatggaaggg 360 aaaatagtgtgactagcatc atcgatttgc ttgctatcat gtttgagcat cattgacatg 420 tgggtgtcgatctaggagac tatgaatcta gcgaatcctc actagtatgc acatgcacaa 480 cgacatcatgcagctatatg tacaacaatg taggaatggt agctctactc ggattggtgg 540 ccgcattcctacattgctag aaaaacacaa acacacacac acacacacac aaacaaacat 600 aggacgctgcaacacctctt ccctggctac cactatcgcc aaatgccaag catggttgtg 660 ttgacctccctccttagctg ccactgtgtc cacttcaaaa tccataacca cgacatcttc 720 ccttcttctacaattgttac tactgttctt gaagaggttc ttgtcctacc gacttactaa 780 agaccaaccattggtcagca gagcatctcg agacttctat accaaagtaa cattgagagg 840 tcaatggcatgtaggaccat cactccgttt gcattgacca atcagagatg agtctgaaca 900 atttcaacccgtaaaaacgg tccacaattt gattcatcta ttggtttgct gtctgcccat 960 ggccctgcatcttgattgtt tatgatcttt cttagttatc tcttgagatt taatctacct 1020 aattgtgtcattttaatcct ttgatttatc ttgattggat tcggtctact ttggtttcac 1080 catcttagccccgatatgat cttggtagcc cttttttcct cgcactaatc attttcgatt 1140 ctcgactccaatctaagttc ccatggtaga tttaggtcaa taagtgaaaa acaatttccc 1200 ttacaatttaatcccctttt gagtaatgac cttcgtactt cctcaaacga tggtttcatg 1260 gataaagatttgatatttgt ggcgcacgga tggggcatgg cttatgaaga aaatgagctc 1320 attggattggtattttctcc tccatgctcc tattggtaaa aatattggtt tagttggata 1380 gtcaggttagaacgggattt aaaaatgtga ctacaactag tcgtttatcg cggttgcaca 1440 tgggtgtcaacgggtgactg cctgtgttgg catcaataat ttagaattag aggtgtcggc 1500 ttaatgctggtagattgtgg atatcctccc atacacgtag gcgtcgatgg cttcatgtgc 1560 gtacgtgccgtcctgcatag gatgggtagc aacggaggaa ggctggcttt gttgacttgt 1620 agcctcactaaggccaactg cctattggca atgaagttgt ttttgtttaa ggagaaacaa 1680 acttttgagctaggcaacat cggtatgtag caggccattc tctaggtcaa tggcacttag 1740 agtcggagcatgccggccgt gcctattaca acatggttat catgtgcatg tggcaagggc 1800 acacattacacgtccccatt gtctaacatc gtagactatg agtttgatag aagaaaccca 1860 actgaaaggtcagtgtcaaa tccattctac ctgtgtcgag aactaaggtt aagcgtttta 1920 tatagattaggacactcaca tacatatgct gtgcatgggg tggtgagagc cattatgtac 1980 cactagtgacatgtgtctac ctagttatat gatctattga ttgaaaattc gactgagata 2040 agaataactatgtggtatat ttaacttggt acaacatata gtggttggaa catcacatta 2100 gacatgccttggcaggaaga aagccgctgt agtttaattt aacatgttaa acaccacata 2160 gtgagtaatcaatctaggct taaaaaaaat aaagacaaca caaacaaaca cagagaacac 2220 ataaacacagagggggcaat caactctgtt gtttgatcta tacttcgtca gagtttgata 2280 tatctctgctctctgcattt taaaactact tatattacaa agcactgtat gttatatagt 2340 aatactttgtagatcacgag tataaaacac agataggtac aatttttttt actcctacgt 2400 atttacgtattaaccggtac aatattattt agcatgcatg tacgcgtgat attagagtgg 2460 agtatactacctacgcatgc tcgggcagta tcggtacctc tatacctacc tatatacaca 2520 taccaggtagagtacagcgg caggtagagt acacagtagt atatggagct tattttatca 2580 tttttattactggagataga agaagaagag aaaggagaga gagagagaaa gagtggaagg 2640 gaaaggcgtgcggggcccac acacgcagca gggcagcgct gcggaggtga ggtgagggca 2700 ggggcaggggcagggcggac tggccaacgc caactccaac tccaactccc gtacaaataa 2760 aatatacaatcctttctttt tcctcctcct cttttccctt ccattccacc cccctctctc 2820 tctcttctccactccaaatc ccttcttacc ctattcccct ccccccgcag cttctcttcc 2880 tcctgcagtactcgccgcca ccaccaccgc gccgccgccg ccggccgcgt tccgagaccc 2940 actcgatcggaatccaccgc ggcgcgcccg cgcgcctgcg tcctcttcct tccccgggag 3000 ccgaccgaccacggcgacca gtcgatctcc ctctccgggc gccaaccgcg tcttagcttc 3060 atcgaatccaccgccccacc ccgcatctcc tcctcctcct ccgacgacga cgactactac 3120 tagtcttctccaataagccc ccctcccgct ccccccgcct gaagaagaag cagcagctag 3180 ctccggggagaggtcgacgg cgcgccgggt agatcgcgcc ccgccccgcc tgcgtcgcgg 3240 ctgtcggagcaaacgcaaac cccccaggta atcaacgaac ttttcctccg ccgcaagaac 3300 agctcccgcggggggtttgg ttttgaccga tttcttcccc cctcccccca aatcgaccca 3360 tccaatttcgcctcgattta cttccgattt ccccactttt ttttcttcct ttcgggttgg 3420 ggggttgcggttttggggga ggagaggggt tcagctcatc cgaagcccca cgttaggtcc 3480 gccccctttccagctgtgcc cctctctcgg gcctcgagct cctcgcctcc atgggaacca 3540 aagcccttatatttcatgtc gcggaagaaa aaaaaatccc gtcttttgcg gggatcctcg 3600 cggctacgtacgagccctag ttaccgcgcg gattttagtt acggcggttt atgcggcccc 3660 tccctctaggttttagatct acccatctct ctctctctct ctctctctct ctctgtgcat 3720 gcatgtgtctatcttagcta tacctgtatt atttggaagg ttaattatgg ttgtgtatat 3780 gtggcgcggtaattaattag tttaattcgc accccctctc tctttgttta tctaggtttt 3840 gggggaatttatttcttgct ataattttgc ccgctcgaat ttctggtgct cttatattcc 3900 atgagctgattgaagtggat atatattgtg cgtgcgtgcg tgctattgct acatcggctt 3960 gacttcttcttgcctactac ttcattaatt tgtttcttct ggtttctgtt tcaggttgtt 4020 ctagcgtgtgcagcggctag ctgattgatt gtcttctgtg atatatccag agctcgtgtt 4080 ttgtggtttgtggtttgtgg tttgtgcttg gattgttgat gtgctaattc gcggcgttac 4140 aagatcactgctggattgat attgagttgt gcctcggctg tgctagctgt gtgttgattc 4200 tctcctcgtcgtggtgatcg atatggagat tgttgcagta gatcaggagg gagctcgtgt 4260 tgttgggacgaactgtatgc ttgctcgtgg tggaactggt gctgtagcgc cagtgttgga 4320 gctgacagcgacgcctcgtc aggatgcagc cgctgaagct ggtgtagacg aaccggcaca 4380 acaccaatgcgagcatttct ccataaggta atcattttct gtatttccaa ttccagtatc 4440 gcgttgtggatgaataatga atcggcatgt catgccatat tgcactgttt gatggaagag 4500 tatgattgatcgtggttttt gcacagtttg ctgttgggac ttatatggtc atctgttttg 4560 tacgatcgtatacactgggt cgacatgctt atgactttgg ttcgatttag gaagtcaata 4620 catccactactagctctata tctagccatg tgaactcatt tatgccatag cacagctagc 4680 aggctagcagcaaaaaatat atataatatt tgcatatatg ttggtgtttc atgtatcttt 4740 atactctacgtacatccatt aatatcttca atgtatgaat ctgagcacat gattgtgagt 4800 gctacacatatgcatgtctg tatgtgtgtt cattaggtgt ttgatcatat ttgtttgtgt 4860 tggggtgcgcatgcatttat tcaggccatg ctgtaggctg tagctagata tttgtgtttg 4920 tatattatttctgttgaaca agctgattac taatgaaatt aacctttttg gggtacactc 4980 atatattgggccctacattt ttgtaatcat ttttcctttg tgctgaggtt cagcataaaa 5040 cttttttatcataagcatgt ttacatccta ggagattctt agaactgatg gtttcttcat 5100 atttgcattatgtttgattt gatagtccat tattatttta agccttttca attgtttaga 5160 gattctagagatgatatata tcaaccatag acttgtcacg ttttggttta atactttcta 5220 gaactaattagattattatt tttgtagttt atcctgtcat gctatttgta ttatctttga 5280 attcaaactgcaatacttag attatcttga aggtcctctt tttctggact gtacaagcta 5340 tgtatgaaatgcctacctcc cagcatcctt tagattatgt agggcctttt ctgagtttat 5400 cagttgtatattgactgaag cacgcaatgt gctatatata tgtgccatgc atctttataa 5460 tgataatcttatttttcttg taccagaggg tatgttgctc ttcttcagaa gaaggatcca 5520 aaattctgctctctatctcg gattttccat gaccagaaaa aatgtgatga acacaaagct 5580 agttcaagcccattttctgt agcaaagttt cgacgatggg attgctcgaa gtgcttggat 5640 aagttgaaaacttcagataa tggaacagca ccaagaactc ttcccgcaaa gcagaatggc 5700 acaagtgatggttgctccat cacatttgtt cggagcactt ttgtgcctgc tagtgttggt 5760 tcccaaaaagtgtctcctag cacacaatca tctcaaggga agaatgctga tagatcaact 5820 cttccaaagagtgtgcaaga aggcaatgac tccaaatgca atgcgccttc tggcaagaat 5880 ggagctgctgaggccaatac tgattcacca atgaaaggta tggtagatgt agagcctttc 5940 aaattcctaagtaggatttt atttaaggta tagaataaac taatgtttgt gtgattttct 6000 cagatttgcaagggccagcc caaaattatg atgtggcagc aaatgtctct gaggacaaca 6060 cttctgttgatgttggggct ttacctgaag ttccccagat tacatggcac atagaagtaa 6120 atggtgcagatcaacctcca tccactccaa aactttctga agtggtcctc aaaagaaatg 6180 aagatgaaaatggaaaaact gaagagactc ttgttgctga gcagtgcaat ttgaccaaag 6240 atcctaacccaatgtctgga aaggaacgtg atcaggttgc tgagcagtgc aatttgacca 6300 aagatccgaaaccagtgtct gggcagaaat gtgagcagat ctgcaatgag ccatgtgaag 6360 aagttgttctcaaaagaagc tccaaatcta agaggaagac ggataagaag ttgatgaaga 6420 agcagcagcacagcaagaaa cgcactgccc aggctgatgt ttcagatgca aagctttgtc 6480 ggagaaagccaaaaaaggtg cggcttctat cagaaattat aaatgctaac caggttgagg 6540 attctagaagtgacgaagtt catcgtgaaa atgccgctga tccctgtgag gatgatagaa 6600 gtaccatcccggtcccgatg gaagtaagca tggatattcc tgttagcaac catacagtgg 6660 gagaagatgggttaaaatca agtaagaaca agacaaaacg caaatactct gatgttgtag 6720 atgatggatcatcacttatg aactggctga atggaaaaaa gaaaagaact ggaagtgtgc 6780 atcacacagttgctcatcca gctgggaatt tgagcaacaa aaaagtgaca cccactgcga 6840 gtactcagcatgatgatgag aatgatactg aaaatggtct tgacacaaat atgcataaga 6900 cagatgtctgtcagcatgta tcagaaatct ccacacagag gtgctcatca aaggggaaaa 6960 cagcgggtttgagtaagggg aaaacacatt cagctgctag taccaaatat ggtggtgaaa 7020 gcaccagaaatggtcagaac atacatgtac tcagcgcaga agatcaatgc cagatggaaa 7080 ccgaaaactctgttctgagt cactcggcaa aggtacgaat tttgtgaatc atgaggaatt 7140 tttgctttttaaattgactg aatcaacatt tatctgtatg aaggaataat attggtgcat 7200 aacaatgttaagaaatatgc atacaatgtt tatttatatg ctttccactg ttcttcttta 7260 cttatgttttgatactcttt ttgtgtgtgc gtgcatgtgt gcatgtgtgt gtgtgtgtgt 7320 gtgtgcgcgcgtgtgtgtgc acgtgcgtgg cgcaatattc ttttttagac tcatattata 7380 gtgattgtaatggactgaca ttttcctcat ttctcatctc aggtttctcc agctgagcat 7440 gatatccaaattatgtctga ccttcatgag cagagtctac ccaagaagaa aaagaagcaa 7500 aaacttgaagtgactcgtga aaaacagacc atgatagatg acatccccat ggatattgtt 7560 gaactgctagctaaaaacca gcatgagagg cagcttatga ctgagactga ttgttctgac 7620 atcaaccgtattcaatccaa gacaactgct gatgatgatt gtgtaatagt agctgccaag 7680 gatggttcagattatgcatc aagtgtgttt gacactaatt cccaacagaa gtccttggca 7740 tcccaaagtacacagaagga gttacagggt catttggcat tgaccacaca agagtctcca 7800 catcctcagaactttcagtc tactcaggaa cagcagacac atttgcggat ggaagaaatg 7860 gtcactattgctgcaagctc accactattt tcacatcatg atgatcagta tattgctgaa 7920 gcaccaactgaacattgggg ccgtaaggac gcaaagaagc taacgtggga gcaatttaag 7980 gccactacaagaaattctcc agcagcaaca tgtggtgctc aatttagacc tggtatccaa 8040 gcagttgacttgacttctac tcatgtcatg ggatcttcca gcaattatgc atctcgccaa 8100 ccagtaattgcgccactgga ccgctatgct gaaagagcgg ttaaccaggt ccatgcaaga 8160 aattttccaagcacaatagc aaccatggaa gcgagtaagt tatgtgatcg gagaaatgct 8220 ggacaagtagtcttgtatcc taaagaatcc atgcctgcga cgcatcttct gagaatgatg 8280 gatccatcaacattagcaag cttccccaac tatggaactt ctagcaggaa ccagatggag 8340 tctcaacttcataattctca gtatgcacat aatcagtaca aaggatcaac cagcacatca 8400 tatggcagtaacctgaatgg aaagattcca ttgacattcg aagacttatc acggcatcag 8460 ctgcatgatctgcacagacc tttacgccca catcctagag ttggtgtgct tggctccttg 8520 ctgcagaaggaaattgcaaa ctggtcggag aactgtggca cacaatctgg ttataagtta 8580 ggagtgtcaacaggaataac atcgcatcag atgaacagaa aggaacattt tgaagccctg 8640 aattctggaatgttttcagc aaaatggaat gcattgcagt tgggttctgt tagctccagt 8700 gcagattttttatcagcgag gaacagcata gctcaatctt ggaccagagg caagggtaaa 8760 atggttcatcccttggatcg gtttgtgaga caggatatct gtataactaa caagaaccca 8820 gctgattttactacaatcag taacgataac gagtatatgg attaccgctg aagcagaaag 8880 tggtgtgcataattcctgaa catttacaat catacatttc atctttatgg cgccaaatag 8940 tcatactgtaagaggagggc tttgctggat ctgctgtaag gtaagttgaa ctttttcttc 9000 ttgcaagtttatcagtttaa gaaaaaagaa tgattactta tgttagcaag gatggttctt 9060 gcaggcttcttgtaagttgt ggatgcccca ttttctggat gggaacctgc cagacagtga 9120 acaagggctttgcaaggtgc agcatccggt ttttgttttg ccagtccaag aaacgtcctc 9180 ctgttactttgtagttgtac tcatactagt gcgcttgttt gtacaaggag aaatgtgtaa 9240 ccttgttgaaaaaatgtctc ccccattttg taattaccat aaggaggttt atagtgttgt 9300 gagctgtgtgtgactgacgg cgagaaatgg ttttgtcggt gttaaggttg aaacgactag 9360 ctctcgttatcaatgtgttg taaacttcta gattgatgtg ttaccttact cttgaagtca 9420 acaccggagaatttacagta cttttttgcc gtgcc 9455

What is claimed is:
 1. A polynucleotide encoding a plant gene capable ofcontrolling a signal transduction system for brassinosteroid hormone,the polynucleotide encoding an amino acid sequence from Met at position1 to Arg at position 1057 of SEQ ID NO: 2 in the SEQUENCE LISTING,including any polynucleotide encoding an amino acid sequence in whichone or more amino acids are deleted, substituted or added to the aminoacid sequence.
 2. A polynucleotide according to claim 1 derived fromrice.
 3. A polynucleotide according to claim 1 as represented by SEQ IDNO: 1 in the SEQUENCE LISTING.